Phycologia (1975) Volume 14 (4), 309-316

The morphology and systematic position of the green algae and

GEORGE F. PAPENFUSS AND Mnsuo CHIHARA

Department of Botany, University of California,Berkeley, California94720, U.S.A. and Department of Botany, Tokyo Kyoiku University, 3-29-1, Otsuka, Bunkyo-ku, Tokyo 112, Japan

The monotypic genera Ernodesmis B0rgesen and Apjohnia Harvey, which in habit resemble each other and which have been regarded as belonging to the by most authors, are shown to be different genera. Neither appears to belong to the Siphonocladaceae. Ernodesmis fits well in the Valoniaceae (Siphonocladales) and Apjohnia is provisionally assigned to the Anadyomenaceae (). Peculiar intracellular rhizoid-like cells are produced at the base of branches in older parts of the thallus of Apjohnia. Comparable structures are also formed in several other green algae, including Ernodesmis and Cladophora rugulosa. They are believed to give mechanical strength at the juncture of adjacent cells, especially at the place of insertion of branches.

Introduction Morphology

From published descriptions and illustrations of B0rgesen (1912, 1913) has fully described and Ernodesmis and Apjohnia, the impression is gained illustrated the development of the thallus; a brief that these two genera of marine green algae may be account will suffice, therefore. The plant starts as congeneric. The present study was undertaken with an erect, clavate, vesicular cell, 10-20 mm long and the hope of resolving this problem. 1·5-2·5 mm in diameter, with basal annulations, attached by a system of septate rhizoids. This cell produces a terminal cluster of five to eight or more Ernodesmis Borgesen one-celled branches, each separated by a concave wall from the parent cell. Each of these primary History of the genus branches produces in turn a similar terminal whorl Ernodesmis was erected by B0rgesen in 1912 (see of branches (Figs 1-3). This process is repeated also B0rgesen, 1913, pp. 66-71) for a species from until six (or more?) orders of branches have been St Croix, Virgin Islands that Kiitzing had described formed. The mature plant attains a height of about in 1849 and illustrated in 1856 (pI. 88) under the 5 cm. Not infrequently, hapteroid rruzoidal cells are name verticil/ata. The plant is widely produced at the proximal end of the cells (see also distributed in the Caribbean (Taylor, 1960) and is B0rgesen, 1912, 1913). Similar rhizoids were called also known from Brazil (Schmidt, 1924), the Canary clamps by Murray & Boodle (1888, p. 276) with Islands (B0rgesen, 1925), several places in West reference to Struvea ramosa. They elongate and Africa (Steen toft, 1967), the west coast of Mexico attach themselves firmly to the distal end of the cell (Dawson, 1961), including the northern part of the below. Gulf of California (Dawson, 1966), Ceylon (Dura i­ The cells are multinucleate and contain numerous, ratnam, 1961), Mauritius (B0rgesen, 1940, 1946) small, polygonal chloroplasts (each provided with a and Kenya (Gerloff, 1960). pyrenoid) joined at the corners to adjacent plastids B0rgesen (1913, 1925) placed Ernodesmis in the to form a reticulum. Siphonocladaceae, in which family it has been The earliest events in the initiation of branches in retained by all authors dealing with it, except Ernodesmis were not observed by us. From slightly Oltmanns (1922), Fritsch (1 935) and Taylor (1960), later stages it is seen, however, that a branch always who placed it in the Valoniaceae. Fritsch and Taylor, has its origin as a lenticular cell formed in the however, did not accept the family Siphonoclada­ terminal region of the parent cell (Figs 1 and 3). ceae. This cell produces a protuberance (Fig. 2) which

309 310 Phyco[ogia, Vol. 14 (4),1975

2 mm

\ 2

FIGS 1-3. Ernodesmis verticil/ato. Stages in the development of lateral branches; note the origin of a branch from a lenticular cell (Fig. I) with a concave basal septum. Papenfuss and Chihara: Ernodesmis and Apjohnia 311 elongates and forms a branch with a concave basal the many species of the genus possess annular septum which has been present since the inception constrictions. of the branch (Figs 1-3).

Apjo/mia Harvey Discussion History of the genus

The method of branch formation in Ernodesmis Apjolmia, with the species A. laetevirens, was is like that described by Papenfuss & Egerod (1957) described by Harvey in 1855 upon material from for pachynema, and also recalls the Australia (Phillip Island, Western Port, Victoria) situation in Valonia (Egerod, 1952). It is unlike the and in 1858 he illustrated it in the first volume of his situation in Siphonoc/adus in which the branches are Phycologia australica (pI. 5). J. Agardh (1887, pp. endogenous in origin as a result of segregative 105-108) reported the species from Tasmania. division of the protoplast of the parent cell (see According to Womersley (1956), A. laetevirens in Egerod, 1952, fig. 2m-o). mainland Australia ranges from Elliston, South Ern.odesmis agrees with Valonia in forming Australia, to Collaroy (near Sydney), New South spherical cytoplasmic masses in some of the cells Wales, whence it was obtained in the drift by May (Fig. 3). These bodies, which were also observed (1946). KUtzing in 1863 (p. 12) described and later by KUtzing (1856) in material of this genus from the (1866, pI. 2) illustrated a plant from New Caledonia Virgin Islands and by B0rgesen (1913, 1940) in (lie des Pins) under the name Struvea scoparia. specimens from the Virgin Islands and Mauritius, Murray & Boodle (1888, pp. 266 and 2 81) merged are frequently present in Valonia, for which they this species in Apjohnia laetevirens, remarking, were first reported by Murray (1893). They have however, that it was not in perfect accord with the been regarded as asexual reproductive structures, latter. Womersley (1956) is inclined to doubt the produced perhaps under unfavourable environ­ conspecificity of these taxa. Denizot (1965) has mental conditions. Their exact function remained rediscovered KUtzing's plant in New Caledonia and uncertain, however, until it was shown by Chihara has discussed and illustrated it under the name (1959) that they can function as aplanospores and A. laetevirens. In habit, the plant from New Cale­ are capable of developing into new plants. donia (Denizot, 1965, pI. [1]) does not appear to Fritsch (1935, p. 424) was of the opinion that correspond exactly to the one from Australia, but 'Ernodesmis, Siphonoc/adus, Struvea, and whether this difference is sufficient to reiain it as form another apparently natural group characterized an autonomous species or to justify its removal to by the annular constrictions on the main axes and another genus cannot be settled without examination sometimes also on the branches'. However, the of the plant. formation of annular constrictions at the bases of The brothers Crouan in 1865 (in Schramm & axes (and at times of the branches) apparently is a Maze, p. 47) described a plant from Guadeloupe feature that has arisen several times in the green Island in the West Indies as Apjohnia tropica. J. alga�, and in most instances it probably is not Agardh in 1887 removed this species to Siphono­ indicative of phylogenetic affinity. Thus, for example, c/adlls Schmitz, where it has remained to the present. Struvea anastomosans, S. tenuis and S. pulcherrima Murray (1891, p. 209) transferred the South (Boodleaceae, Siphonocladales) lack annular con­ African Cladophora 1"1Iguiosa Martens (1866) to strictions in the stipe whereas the other species of Apjohnia. Although there are features of agreement the genus possess them (Egerod, 1952, p. 361); between A. laetevirens and C. ruglllosa,the evidence some species of Caulerpa (Caulerpales), e.g. the favours keeping C. rugulosa in the genus Cladophora. South African C. /ili/ormis and C. holmesiana, Wille (1910, p. 111) reduced the monotypic possess annular constrictions in the stipe whereas Rudicularia Heydrich (J 903), based on material the great majority of species of the genus lack them; from the Ryukyu Islands, to a synonym of Apjohnia in the Cladophorales, Chaetomorpha antennina and made the combination A. penicillata (Heydrich). possesses annular constrictions but all the other RlIdiclllaria probably has little, if anything, in com­ species of the genus, to our knowledge, lack them; mon with Apjohnia. It may be congeneric with in Cladophora itself, as far as we are aware, only Tydemania Weber-van Bosse but for the present its C. rugulosa, C. wrightiana and C. prolt/era among identity must be regarded as uncertain. (See Papen- 312 Phy'!;Vlogia, Vol.14 (4),1975 fuss, 1953, p. 30, for a brief discussion of Rudicu­ dendroid tuft to 15 cm high (Harvey, 1858). It is laria.) attached by a system of branched, sparsely septate Harvey (1855), J. Agardh (1887), Murray (1895), rhizoids. The plant starts as an erect, club-shaped Wille (1890, 1910) and Printz (1927) placed Apjohnia vesicular cell, which produces rhizoids at its proximal in the Valoniaceae whereas Oltmanns (1904), end and at its summit a dome-shaped outgrowth Feldmann (1938) and Womersley (1956) placed it in which will become an axial cell. After the dome has the Siphonocladaceae. B0rgesen (1913) and West been formed, a whorl of conical outgrowths with (1916) treated the Siphonocladaceae as a subfamily obtuse apices is produced round the central axial of the Valoniaceae and referred Apjohnia to it. cell (Figs 4 and 5). All of these outgrowths elongate Several authors (Wille, 1890; B0rgesen, 191 2 ; considerably and are ultimately separated by basal Printz, 1927) considered Apjohnia to be a nonseptate septa from the lower primary cell, which functions alga, an erroneous belief that caused Setchell (1929) as a stalk cell. The cells surrounding the axial cell to suspect that the genus should be placed in develop into a whorl ofJatera\ branches. This process the Codiales as the representative of a family of axial cell and lateral branch formation is repeated Apjohniaceae. a number of times and the primary laterals also produce whorls of laterals of a second order; as many as five or more orders of whorled lateral Morphology branches may be formed. As a rule only two, more The thallus of Apjohnia is in the form of a stipitate, or less opposite, branches are produced at each

4

6 7 8 9

FIGS 4-9. Ap johnia laetevirens. Figs 4 and 5. Stages in the development of whorled lateral branches; note that in these early stages of branch formation basal septa are lacking (later, cross walls are produced). Figs 6-9. Terminal region of thallus, showing the formation of only two, more or less opposite, branches at each node in contrast to the whorl of branches at each node in the lower part of the plant. Papenfuss and Chihara: Ernodesmis and Apjohnia 313

node in the terminal region of the thallus (Figs 6-9) D. versluysii, as illustrated by Egerod (1952, Fig. in contrast to the whorl of branches at each node in 2i-k). the lower part of the plant. Structures comparable to those in Struvea ramosa The stalk cell in the mature plant may attain a were observed by Kuckuck (1907, p. 181), B0rgesen length of 5 cm and a diameter of 4 mm. Harvey (1912, Fig. 11c; 1913, Fig. 53c) and by us, as pointed (1858) was of the opinion that the stalk cell becomes out above, in Ernodesmis verticillata. As in S. calcified but this is not so; the calcification is owing ramosa, they are on the outside of the cells. to the presence of epiphytic crustose coralline algae. Somewhat analogous bodies have been observed The upper segments are 1-2 cm long and 1-2 mm by Chihara (1960) and Sakai (1960, 1964) in Clado­ in diameter. Annular constrictions develop at the phora wrightiana Harvey. However, in this species bases of the stalk cell and all the mature segment cells the rhizoidal structures (or the 'non-septate ... of the axis and lateral branches. Dawes (1969, p. 81) protoplasmic protuberances' as Sakai, 1960, p. 4, observed that 'cross walls were common across calls them) do not become separated by a wall from annular constrictions found at tbe base of all the protoplast of the parent cell. At the juncture of filaments and at the junction of rhizoids and the adjacent cells from four to ten of these short, stalk cell'. rhizoid-like processes may be formed at the lower We found no evidence of segregative cell division end of the upper of two adjoining cells. The processes in the production of axial and branch cells, an grow downward through the septum between the two observation which confirms the statement by Dawes cells arid surround the upper end of the protoplast (1969, p. 81) that 'the development of the cross walls of the lower cell (see Sakai's, 1960, excellent figures in the segments and filaments apparently proceeds of them). centripetally from the side walls toward the center, Similar rhizoid-like processes occur in South in the manner described for the Cladophorales African specimens of Cladophora rugulosa Martens (Fritsch, 1935)'. (Figs 12 and 13), a species which shares certain A striking feature of Apjohnia is the formation of other features with C. wrightiana, and which Murray small, intracellular, rhizoid-like cells at the base of (1891), as pointed out above, regarded as belonging branches in older parts of the thallus. These small to Apjohnia. Sakai, in his monograph of the Japanese cells, four to ten in number, are produced at the species of Cladophora, makes no mention of such base of the parent cell and grow downward into the processes in Japanese specimens of C. rugulosa. If cross wall. They may penetrate into the upper part they should be absent in Japanese material, grounds of the segment cell below, where their arms attach may exist for regarding the Japanese plant as themselves very firmly to the inner surface of the representative of a different species. * wall of that cell (Figs 10 and 11). The rhizoidal We also examined British, Adriatic and Israeli processes of these cells apparently never penetrate material of Cladophora proU/era (Roth) Kiitzing, to the outer surface of the cells. which shares certain features with C. wrightiana These peculiar structures were also seen in and c: rugulosa, but did not see any endogenous, Apjohnia by Dawes (1969, p. 81), who called them rhizoid-like, protoplasmic processes in this species. cyst-like cells. Similar organs, except that they are Nor are they reported for it by van den Hoek (1963) produced on the outer surface of the cells, were in his monograph of tbe European species of observed and illustrated for the first time in Struvea Cladophora. ramosa Dickie by Murray & Boodle (1888, figs 3g, h) All these organs probably perform an important who (p. 276) called them clamps and remarked about function in giving the plants in question mechanical them as follows: 'They have evidently been produced strength at the juncture of adjoining cells, especially by the formation of a small wart-like outgrowth at the base of branches. In Struvea ramosa, Murray from the base of a branch just above its basal wall. & Boodle (1888) observed them in the basal region The outgrowth then grew downwards parallel to the

filament, producing a neck-like prolongation which * Martens (1866) when describing Cladophora rllglliosa curved inwards and came into contact with the referred to it specimens from both Port Natal, South Africa surface of the filament below the transverse wall, and Yokohama, Japan. Papenfuss (1943) regarded Port and then threw out a fan-shaped mass of branched Natal as the type locality. Subsequently the first author saw an authentic South African Martens specimen and rootlets on the wall'. These organs are reminiscent accompanying diagnosis in the Rijksherbarium, Leiden. of those in , especially those in This specimen is here designated lectotype of this species. 100 pm I

50 pm 0·5 mm

12 11

FIGS 10 and 11. Apjohnia laetevirens. Small, intracellular rhizoid-like cells formed at the base of branches in older parts of the thallus. FIGS 12 and 13. Cladophora rllglliosa. Rhizoid-like processes, which do not become separated by a wall from the protoplast of the parent cell, formed at the lower end of the upper of two contiguous cells. Papenfuss and Chihara: Ernodesmis and Apjohnia 315 of the frond where the main branches are formed. Acknowledgments In Cladophora wrighfiana, Sakai (1960, 1964) saw them at the places where branches were inserted. The authors are indebted to Dr L.R. Almodovar In Apjohnia laetevirens we observed them only at the for sending them formalin-preserved material of base of branches which usually themselves bore Ernodesmis from Puerto Rico and to Dr H.B.S. branches, and in C. rugulosa we saw them only in Womersley for providing them with formalin­ the lower part of the thallus. preserved material of Apjohnia from South Australia. Herbarium specimens of Apjohnia from South Australia, previously given to the University of Discussion California Herbarium by Dr Womersley, were also used in our study. The work was supported by a The taxonomic position of Apjohnia is problema­ grant (0-24302) from the U.S. National Science tical. The genus has been placed in the Siphono­ Foundation to the first author, and was done in c1adaceae but we have seen nothing in it suggestive Berkeley during 1962-1963, when the second author of segregative division, the method of cell division was on leave of absence from Shimoda Marine characteristic of Siphonocla-lis (Egerod, 1952, Fig. Biological Station. 2m-o; Christensen, 1966, Fig. 71a, b, d). In fact, we have not observed any feature that would justify retaining it in any of the families of the Siphono­ References c1adales. Itshares features with members of both the Cladophoraceae, especially Cladophora rugu/osa AGARDH, J.G. (1887) Till algernes systematik ... (Afd. 5). Lunds Univ. Arsskr.,23 (Afd. 3, No. 2), 174+ [6] pp., 5 pIs. and C. wrightiana, and the Anadyomenaceae, B0RGESEN, F. (1912) Some Chlorophyceae from the Danish families in the Cladophorales. On the basis of the West Indies. ll. BOI. Tidsskr.,32, 24 1-273, 17 figs. resemblance of the terminal parts of mature thalli, B0RGESEN, F. (\913) The marine algae of the Danish West Indies. Part l. Chlorophyceae. Dal1sk. Bol. Arkiv, 1 (4), where only two lateral branches, instead of a whorl, [2]+ 158+ [2] pp., 126 figs, 2 maps. are produced at a node, to the terminal parts of B0RGESEN, F. (1925) Marine algae from the Canary Islands ... l. Chlorophyceae. K. Dal1ske Videl1sk. Selsk. Bioi. thalli of Willeella (Chihara, 1965, Figs 4A, 5) of the Medd.,5 (3), 123 pp., 39 figs. Anadyomenaceae, we provisionally assign the genus B0RGESEN, F. (]940) Some marine algae from Mauritius. I. to this family. Perhaps it should be placed in the Chlorophyceae. K. Danske Videl1sk. Selsk. Bioi. Medd., 15 (4), 81 pp., 26 figs, 3 pIs. Cladophoraceae. B0RGESEN, F. (1 946) Some marine algae from Mauritius. With respect to the systematic position of Apjohnia, An additional list of species to part I Chlorophyceae. it is to be noted that Oltmanns (1922), Feldmann K. Danske Vidensk. Selsk. Bioi. Medd., 20 (6), 64 pp., 27 figs. (1938), Chapman (1954), and Christensen (1966) do CHAPMAN, V.J. (1954) The Siphonocladales. Bull. Torrey Bol. not regard the Cladophorales and Siphonocladales Club, 81, 76-82, 4 figs. as separate orders, a view accepted also by Dawes CHIHARA, M. (1959) Studies on tbe life-history of the green algae in the warm seas around Japan (9). Supplementary (1969, 1974); they combine these two groups under note on the life-history of Valonia macrophysa Klitz. J. the name Siphonocladales. However, the peculiar Jap. Bot., 34, 257-266, 5 figs. CHJHARA, M. (1960) Studies on the life-history of the green method of cell division (segregative division) algae in the warm seas around Japan (10). On the life­ characteristic of some members of the Siphono­ history of Cladophora wrighliana Harvey. J. Jap. BOI., 35, c1adales, especially Siphonoc/adus, Dictyosphaeria 1-11, 4 figs. CHIHARA, M. (1965) The life history and of and Struvea, set these algae sharply apart from Willeella japol1ica. Bul/. Nal. Sc. Mus. Tokyo, 8, 355-368, members of the Cladophorales. The endogenously 6 figs. formed lenticular cells and the aplanospores of CHRISTENSEN, T. (1966) Syslematisk BOlanik. Nr. 2, Alger. Munksgaard, Copenhagen, 180 pp., 76 figs. certain Valoniaceae conceivably represent aspects DAWES, c.J. (1969) A study of the ultrastructure of a green of segregative division. An ultrastructural study of alga, Apjohl1ia laelevirens Harvey with emphasis on cell wall structure. Phycologia,8, 77-84, 11 figs. the method of formation and ultimate structure of DAWES, c.J. (1974) Marine Algae of the West Coast 0 these various cell types would be instructive. If Florida. University of Miami Press, Coral Gables, Florida, future studies show that the Cladophorales and xvii + 20 I pp., 82 figs, 3 tables. DAWSON, E.Y. (1961) A guide to the literature and distribu­ Siphonocladales (Blackman et Tansley) Oltmanns tions of Pacificbenthic algae from Alaska to the Galapagos (J 904) indeed should be combined, the name that Islands. Pac. Sc.,15, 370-46 1. has the right of priority is Cladophorales Haeckel DAWSON, E.Y. (1966) Marine Algae in Ihe Vicinity of Puerto Peiiasco, Sonora, Mexico. University of Arizona, Tucson, (1894, pp. 302, 305). iii+57 pp., map. 316 Phyc% gia, Vol. 14 (4), 1975

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(Accepted 6 June 1975)